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\3 Biodiversity 

Bulletin of the Torrey Botanical Club. 

New York Torrey Botanical Club,1 870-1 996 

V.35 (1908): 

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Vol. 35 No 4 




APRIL, 1908 
Hepaticae of Puerto Rico 



Alexander William Evans 
(with plates 6-8) 


The genus Brachial ejeunea includes between twenty and thirty 
recognized species. About half of these are American, the others 
being found in eastern Asia or among the islands of the Pacific. 
No species has as yet been reported from Africa. Although essen- 
tially tropical in its distribution, the genus reaches its northern 
limits in Florida and japan and its southern in Australia and Pata- 
gonia. Since the first species described by Spruce is B. laxifolia 
(Tayl.) SchifTn., this may be considered the type of the genus. It 
was originally described from specimens collected by Jameson in 
Ecuador but is now known also from Bolivia. 

With scarcely an exception the species of BracJiiohjamca are 
found on trees or on rotten logs. In many cases they grow mixed 
with other Lejeuneae or with Frullaniae but they sometimes form 
extensive mats by themselves. The plants are usually more or 
less pigmented and often appear very dark brown or nearly black, 
with little or no indication of glossiness. The prostrate stems 
cling closely to the substratum by means of numerous rhizoids, 
which take their origin from rudimentary discs at the bases of the 
underleaves. The branching is at first irregularly pinnate, but the 
female plants after flowering usually exhibit a false dichotomy 
very much as in Marchcsinia. 

The leaves are imbricated and sometimes densely so. In dry 

[The Bulletin for March, 1908 (35 : 97-154, A- /-J)» «"* is sued 20 Ap 1908.] 


156 Evans: Hepaticae of Puerto Rico 

plants the lobes are suberect and wrapped around the stem, but as 
soon as they become moistened they spread widely (plate 6, fig- 
ure i) and assume a more or less squarrose position. The change 
in appearance is very similar to what is found in the genus Mastt- 
gotejeunea. The lobes are approximately ovate in outline and vary 
at the apex from rounded to apiculate or acute. The margin is 
usually entire, but, in certain species, tends to be slightly serrulate 
in the neighborhood of a female inflorescence. The lobule is 
relatively large and consists of two distinct portions approximately 
equal in size {figures I, 2). One of these occupies the carinal 
region and takes part in the formation of a conspicuous water-sac, 
usually about half as long as the lobe. The other is bounded by 
the free margin and is closely appressed to the lobe except at the 
apical sinus where an opening into the sac is to be observed. The 
appressed portion of the free margin bears a series of teeth from 
three to ten in number. These teeth vary greatly in size and in 
form (figures 7-9), but in the majority of cases each tooth is sev- 
eral cells long and so strongly curved toward the lobe that it can- 
not be straightened out even by pressure on the cover-glass (fig- 
ure 1 2). The outermost tooth, which represents the apex of the 
lobule, varies from a single projecting cell to a broad and rounded 
projection (figures 10, 11). The hyaline papilla is borne at the 
proximal base of this tooth but is displaced to the inner surface of 
the lobule, usually appearing two or three cells from the margin. 
Beyond the apical tooth the long and shallow sinus extends, form- 
ing a very acute angle with the outer part of the keel. The latter 
is more or less arched near the base but tends to be incurved where 
it meets the postical margin of the lobe. The leaf-cells have thm 
walls but distinct trigones (figure 6). These are usually triangu- 
lar in outline, two of the sides being convex and one concave. 
The thin places between the trigones are relatively wide and rarely 
develop intermediate thickenings. 

The undivided underleaves are rotund to reniform in outline 
and are attached by an arched line (figure i). In certain species 
they are distinctly a uricu I ate at the base (figure 1 4); in others 
they arc rounded or even cuneate. The margins are sometimes 
plane and sometimes more or less re volute ; they vary from entire 
to irregularly sinuate but are never distinctly toothed. 

Evans : Hepaticae of Puerto Rico ' 157 

In the majority of the species two types of branching are to be 
observed, very much as in the genus Bryopteris and in certain 
other genera of the Lejeuneae Holostipae* In at least one species, 
however, all of the branches seem to conform to the Lcjeunea type, 
being borne behind leaves with lobules- The Fmllania type of 
branching, in which the subtending leaves fail to develop lobules, 
is largely restricted to robust vegetative axes. In a branch of this 
character the subtending leaf is partly inserted on the main axis 
and partly on the branch, the postical base being slightly revolute 
(figures 2, 13). The first underleaf is usually distinctly bilobed 
and is displaced in such a way that the branch seems to arise from its 
axil. It embraces the base of the branch and partially enwraps 
the postical base of the subtending leaf. The first leaf is of small 
size and complicate-bilobed, but the lobule is explanate and rounded 
at the apex. The succeeding leaves and underl eaves are normal 
in appearance. The Lejewiea type of branching, even where it 
does not occur in the vegetative portion of a plant, is almost 
invariably associated with subfloral innovations. An exception to 
this condition, however, is found in the remarkable B. sandvi- 
censis (Gottsche) Evans, of eastern Asia and the Hawaiian Islands. 
In this species the bract behind which a subfloral innovation arises 
is wholly destitute of a lobule and is partially inserted on the inno- 
vation itself. 

The inflorescence in Brackiolejcunea seems to be fairly constant 
for a given species and may be dioicous, autoicous, or paroicous. 
The female branch varies greatly in length but is usually distinctly 
elongated. Subfloral innovations are invariably present and usu- 
ally occur in pairs (figures 4, 5) ; in rarer cases only one innova- 
tion is developed. The innovations are often floriferous and give rise 
to the false dichotomy which is characteristic of the genus. The 
bracts are scarcely complicate and the lobe is usually more pointed 
than in the leaves. In many species a wing is developed at the 
base of the keel (figures 4, 5)- The bracteole is free or nearly so 
and varies at the apex from rounded to retuse or shortly bilobed. 
The perianth scarcely projects beyond the bracts unless the basal 
portion elongates with the development jrf^sp orophyte. It is 

^ — -^ ^M M I — ~ ^^^" 


* See Evans, Hull. Torrey Club 34 : 559- l 9°$- 
t Trans. Conn. Acad. 10: 419. 1900. 

158 Evans: Hepaticae of Puerto Rico 

approximately obovoid in shape and rounded to truncate at the 
apex, with a short beak. It is sometimes terete and sometimes more 
or less compressed, but even in the latter case the lateral keels are 
never sharp. The postical surface bears from two to five keels, and 
the antical surface is either plane or provided with three or four keels 
(figures 21, 22). All of the keels are rounded and are never 
winged or toothed in any way. In pluriplicate perianths they are 
separated from one another by deep grooves, and there is no dif- 
ference, except in position, between the lateral keels and the others. 
In autoicous and dioicous species the male spikes are long and 
often proliferate at the apex. The bracts are imbricated and tend 
to be smaller than ordinary leaves, but their lobules are relatively 
larger and more strongly inflated. Even here the free margin of 
the lobule is usually more or less toothed. The antheridia occur in 
pairs, and the bracteoles are found throughout the entire length of 
the spike. In paroicous species the bracts are much fewer and are 
essentially like normal leaves ; they differ also from the bracts just 
described in bearing the antheridia singly. 

The present genus is in most respects clearly defined. The 
squarrose leaves, the large lobules with teeth along the free mar- 
gin, the subfloral innovations usually occurring in pairs, and the 
plicate perianth with unarmed keels afford an excellent combina- 
tion of generic characters. Certain species, however, as Spruce 
has already pointed out, show a relationship with MarcJmima, 
while others are even more closely allied to the genus Ptychocoleus 
as restricted in the present paper. In most species of Marchesinia 
the free margin of the lobule is also denticulate and the sub- 
floral innovations occur in pairs, but the leaves are not squarrose 
and their lobes are relatively smaller and very different in appear- 
ance from those found in Brachiolejeunca. The perianth, more- 
over, is distinctly compressed and plane on both antical and postical 
surfaces. The characters which separate Brachiolejeunea from 
Ptychocoleus will be considered under the latter genus. 

Two species of Brachiolejeunca, B. densifolia (Raddi) comb, 
nov. jndif. corticalis (Lehm. & Lindenb.) Schiffn., have been re- 

f„^ l^t T C,e * 1S Commonl y known as B. bicohr (Nees) Schiff,,., in spite of the 
.nnliH f """•"*' demi f° lia Ra <Wi and Jungtrmanma ticolor Nees have long been 
cohered S ynony ms . Trevisan restored Raddi's specific name, but has not been fol- 

Evans: Hepaticae of Puerto Rico 159 

ported from the West Indies. Neither of these has yet been found 
in Puerto Rico, but a third species occurs in recent collections from 
the island. It is apparently undescribed and may be characterized 
as follows : 

Brachiolejeunea insularis sp, nov. 

Dull-yellowish or brownish-green, sometimes almost black, 
scattered or growing in depressed mats : stems 0.25 mm. in diam- 
eter, sparingly pinnate, the branches obliquely to widely spread- 
ing, similar to the stem but with somewhat smaller leaves, never 
microphyllous : leaves closely imbricated, the lobes slightly falcate, 
ovate, 1.2-1,7 mm. long, 0.85-1.2 mm. wide, rounded to subcordate 
at the base and rounded or very obtuse at the apex, margin entire, 
strongly outwardly curved from the antical base to the apex; lobule 
ovate-triangular in outline, 0,85-1 mm. long, 0,35-0.45 mm. wide, 
the inflated portion forming a conical water-sac about half as long 
as the lobe, keel nearly straight from a more or less arched base, 
usually forming a continuous line with the postical margin of lobe, 
free margin rounded to cordate at the base, usually bearing from 
eight to ten more or less distinct teeth, those normally developed 
two or three cells lone, one or two cells wide at the base and curved 
inward toward the lobe, apical tooth very variable ; cells of lobe 
more or less convex, averaging 14 [t at the margin, 28 x 22 fit in the 
middle and 35 x 28/^ at the base, intermediate thickenings infre- 
quent, oval : underleaves loosely imbricated, plane, broadly orbic- 
ular, 0.6-0.7 mm - I° ll g» o<75~0-85 mm. wide, distinctly auriculate 
at the base with crenulate auricles, margin otherwise entire or ir- 
regularly sinuate, apex broad, truncate to retuse : inflorescence 
paroicous : 9 inflorescence borne on a long branch and innovating 
on both sides ; bracts erect-spreading, indistinctly complicate and 
unequally bifid, the lobe ovate to oblong, 1 .4 mm. long, 0.85 mm. 
wide, rounded to obtusely pointed at the apex, margin more or less 
sinuate and crispate but not toothed, lobule adnate to lobe for 
greater part of its length, ovate to oblong, 0.85 mm. long, 0.35 
mm- wide, apex mostly acute, rarely blunt, margin mostly entire but 
rarely with a tooth near the apex, wing broad, approximately semi- 
circular, entire, usually adnate for its whole length ; bracteole 

lowed by subsequent authors. The synonymy of the species is as follows : Fruilanoides 
dtmifoha Raddi, Mem. Soc. liai. Modena Fis, 19 : 3 8 - l82 3 J a0: P L 2 < /■ 5- 
1829, Jun^trmannia bicohr Nees, in Martius, Fl, Bras. I 1 : 349. 1833, Lejtunta 
bicohr Nees, in Montagne, Flor- Boliv. 66; d'Orbigny, Voy, dans 1'Amir, Merid. 7 1 . 
*839- Phragmicoma bicohr Nees, in G. L, & N. Syn. Hep. 294. 1845. Ptycko- 
toltus dcmifolius Trevis. Mem. 1st. Lomb, III. 4: 405. 1877. Lejtunea {Brachio- 
Lejtunta} bicolor Spruce, Hep. Amaz. et. And. 131. 1 884. Brachioitjtunea 
*wfor Schiffh. in Engler & Prantl, Nat. Pflanzenfam. i 3 \ 128. 1895. 

160 Evans: Hepaticae of Puerto Rico 

oblong to obovate, 1.2 mm. long, 0.85 mm. wide, plane or nearly so, 
margin entire or irregularly sinuate, apex broad, truncate or sub- 
retuse ; perianth slightly exserted beyond the bracts, oblong- obo- 
void, 1.5 mm. long, 0.85 mm. wide, rounded to truncate at the apex, 
slightly or not at all compressed, mostly ten-keeled, the keels ex- 
tending to below the middle, rounded and separated by deep 
grooves : c? bracts in one or two pairs below the involucre, essen- 
tially like the ordinary leaves : mature sporophyte not seen. 
(Plate 6.) 

On trees and logs. Near Mayaguez, Heller {4.4.63a), Near 
Cayey, Evans (p/). Mount Morales, Utuado, Howe {465). The 
writer's specimens from near Cayey may be designated the type. 
The species has also been collected in Cuba, Wright, Undenvood & 
Earle, and in Jamaica, Underwood \ Evans. 

B. insularis is closely related to B. densifolia, and the two 
species have been more or less confused. B. dcnsifolia was origi- 
nally collected in Brazil, where it seems to be abundant and widely 
distributed. Its range extends also along the chain of the Andes 
from Colombia to Bolivia. In North America it has been recorded 
from Mexico by Gottsche and from the island of St. Vincent by 
Spruce. It resembles B. insularis in general habit, in its auriculate 
underleaves and in its pluriplicate perianth. It is markedly distinct, 
however, in its dioicous inflorescence, the male inflorescences form- 
ing long spikes with closely crowded bracts. It differs also in its 
greater size, in its more sharply pointed leaves, in the fewer and 
smaller teeth along the free margin of the lobule and in its revo- 
lute underleaves. Even in B. densifolia the leaves are sometimes 
rounded as in B. insularis, but this is an exceptional condition and 
is usually associated with incomplete development. 

B. corticalis, which is surely to be expected in Puerto Rico, is 
considerably smaller than B. insularis, and the teeth along the 
free margins of the lobules are fewer and simpler, each tooth con- 
sisting usually of a single projecting cell. The species is further 
characterized by its dioicous inflorescence, by the more distinct 
bracteal lobules, ligulate-Ianceolate in outline and rounded at the 
apex, and by the more strongly compressed perianth with fewer 
and more irregular keels. Another close ally is B. chinantlana 
(Gottsche) Schiftn.,* at present known from Mexico and Ecuador. 

* Hedwigia 33 : 180. //. 9 .f. J2 - 40 . ,8 94 . The spec i es i s based on Phragmi 
coma buotor var. thinantlana Gottsche, Mex. Leverm. 172. 1863. 

Evans: Hepaticae of Puerto Rico 161 

This species agrees with B. insularis in its inflorescence, but the 
lobes of its leaves and bracts are sharply pointed. The bracteole 
is also said to be emarginate or even bifid, but the figure of 

Schiffner does not show this condition clearly. 


The history of Trevisan's genus Ptychocoleus has already been 
noted by the writer in another connection.* It is based on 
Phragmicoma, § Ptychanihoides of the Synopsis Hepaticarum, and 
most of its species would be included in the genera Acrolejeunea, 
Brachiolejeunea or Mastigolejeunca, as these are at present under- 
stood. The first species listed by Trevisan is P. aulacophorus 
(Mont.) Trevis. This was originally described by Montagne.f 
under the name Phragmicoma aulacophora, from specimens col- 
lected in the Mangareva or Gambier Islands, of Spencer Gulf, 
South Australia, but is now known to have a wide range extend- 
ing into both Asia and Africa. Ptychocoleus aulacophorus would 
naturally be selected as the type of the genus. This species, how- 
ever, is placed by Stephani in Acrolejeunea, and it therefore seems 
justifiable, in restoring the name Ptychocoleus, to apply it to the 
genus Acrolejeunea as defined by Spruce and by Schiffner. 

In this restricted sense Ptychocoleus would include between 
forty and fifty recognized species, all of which are tropical in their 
distribution. It attains its best development in southeastern Asia 
and the islands of the Pacific but is also well represented in Africa. 
In America five species are at present known, three of which have 
been reported from the West Indies. Only one species, however, 
P. poly car pus (Nees) Trevis., has been collected in Puerto Rico. 
As in the preceding genus, practically all the species of Ptycho- 
coleus grow on trees or on rotten logs. 

The genus is characterized by an elongated female branch 
without innovations and by a plicate perianth with smooth keels 
(plate 7, figure 2), the number varying from four to ten in dif- 
ferent species. The absence of subfloral innovations will at once 
distinguish it from Brachiolejeunea, but the two genera agree so 

* Bull. Torrey Club 34 : 543- l 9 oS - „ . - ., 

t Ann. Sci. Nat. Bot. II. ig : 259. 1S43. Voy. au FOle Sud, Bot. I i, 

/y./ /. 1845. 

1^2 Evans : Hepaticae of Puerto Rico 

closely in the characters derived from the lobes of the leaves, 
from the underleaves, and from the bracts and perianths that it 
would be superfluous to describe these organs in detail for Pty- 
chocoleus. There are also no essential differences in color, in 
general habit, or in cell-structure. The lobules of the leaves, how- 
ever, are much more variable in Ptychocoteus than in Brachiolejeunea, 
especially with respect to the number of teeth on the free margin. 
Certain species develop as many as four or five teeth, others only 
two (figure 5), while in still others the margin is quite entire 
except for the apical tooth. In the position of the hyaline papilla 
the genus agrees with Brachiolgauiea, except for the fact that it is 
sometimes so much displaced that it is difficult to determine 
whether it is distal or proximal to the apical tooth, while on the 
other hand it may arise directly from a marginal cell. The position, 
however, is usually constant for a given species. The branching, 
so far as observed, always conforms to the Lejeunea type. 

The genus shows but little relationship with other Lejeuneae 
which lack subfloral innovations. In Lophokjeunea the keels of 
the perianth are winged and variously toothed or laciniate, the 
lobules are built up on a different plan, and the local thickenings 
in the cell-walls are different. In Caudalejeunea the perianth is 
trigonous, its keels are normally winged and toothed, the plants 
are different in habit, and the leaf-cells show very numerous inter- 
mediate thickenings. In Bryopteris the female branch is much 
shorter, the perianth is again trigonous, the leaves and under- 
leaves are sharply toothed, and the whole habit, general appear- 
ance, and cell -structure are different. 

Ptvchocoleus polycarpus (Nees) Trevis. 
Jungcrmannia polycarpa Nees, in Martius, Fl. Bras V • 350. 

Phragmicoma polycarpa Nees, in G. L. & N. Syn. Hep. 295. 1845. 
Lejatuea domingensis Tayl. Lond. Jour. Bot. 5 : 389. 1846. 

Ptychocoleus polycarpus Trevis. Mem. 1st. Lomb. III. 4: 405. 

Lijainea (Aero -Lejeunea) domingensis and polycarpa Spruce, Hep. 
Amaz. et And. 116. 1884. 

>jettnea polycarpa Schiffn. in Engler & p rant l ( Nat. Pflanzen- 
fam. i 3 : 128. ' 1895. 

Evans : Hepaticae of Puerto Rico 163 

Olive-green, not glossy, scattered or growing in depressed 
mats: stems 0.17 mm. in diameter, irregularly pinnate, the 
branches widely spreading, occasionally microphyllous : leaves 
densely imbricated, the lobe suberect and convolute when dry, 
widely spreading and squarrose when moist, more or less convex, 
broadly ovate, slightly falcate, 1-1.2 mm. long, 0.85-0.95 mm. 
wide, arching partially or wholly across axis, rounded at base, 
rounded to very obtuse at the apex, margin entire, strongly out- 
wardly curved from the antical base to the apex ; lobule ovate, 
0.6 mm. long, 0.35 mm. wide, the inflated portion occupying a 
little more than the carinal half, keel arched, forming an obtuse or 
rounded angle with the postical margin of lobe, free margin 
straight or a little curved, often bearing a tooth one or two cells 
long in the vicinity of the apical tooth, the latter mostly two cells 
long and one and two cells wide at the base, slightly outwardly 
curved, hyaline papilla borne on a marginal cell but slightly dis- 
placed to the inner surface of the lobule ; cells of lobe plane or 
nearly so, averaging l$p at the margin, 2} x 1$ ft in the middle 
and 35 x 18// at the base, intermediate thickenings occasional, 
oval or circular: underleaves loosely imbricated, plane or slightly 
revolute at the apex, broadly orbicular-obovate, 0.6 mm. long, 
O.75 mm. wide, cuneate toward base and attached by an arched 
line, broad and truncate at the apex, margin entire or vaguely 
angular-denticulate : inflorescence autoicous or polyoicous : $ 
inflorescence borne on an ascending and more or less elongated 
branch ; bracts widely spreading, complicate with a rounded keel 
and shortly and unequally bifid, the lobe ovate, 1.6 mm long, 0.85 
mm. wide, acute and apiculate, the apex usually tipped with a row 
of two or three cells, margin entire, lobule acuminate, otherwise 
similar to the lobe, 1.4 mm. long, 0.4 mm. wide; bracteole free, 
oblong or ovate, 1.4 mm. long, 0.85 mm. wide, undivided and 
acute or shortly bidentate at the apex with sharp or blunt teeth, 
margin otherwise entire ; perianth about one third exserted, 
obovoid, 1.5 mm. long, O.85 mm. wide, slightly compressed and 
cuneate toward the base, truncate to retuse at the apex, antical face 
plane or nearly so, postical face with two confluent rounded keels 
often extending beyond the beak : cf inflorescence terminating a 
more or less elongated simple branch, not proliferating; bracts 
mostly in eight to twelve pairs, shortly and unequally bifid with a 
strongly arched keel, lobe obliquely spreading, more or less 
squarrose, rounded, lobule suberect, mostly acute; antheridia 
borne singly ; bracteoles extending the whole length of the spike, 
similar to the underleaves but more often revolute at the apex : 
capsule about 0.5 mm. in diameter, walls of inner layer of cells 
thickened on the inside with a few large circular or oval pits ; 

164 Evans : Hepatic ak of Puerto Rico 

spores green, spherical or ellipsoidal, 35 /1 in diameter, minutely 
verruculose and with circles of indistinct radiating ridges ; elaters 
about 10 « in diameter, sometimes bispiral in the middle. (Plate 
7, figures 1— 1 1.) 

On trees. Between Cayey and Caguas, Howe (1411 p- P-> 
1414). Type locality, Brazil, Martins ; since found by numerous 
collectors. Known also from Mexico, Liebman, and from the fol- 
lowing islands of the West Indies: Cuba, Wright; Santo Do- 
mingo, Pcrsoon (the type locality of Lejeuttea domingensis); Ja- 
maica, Harris, Evans. 

The species is not well described in the Synopsis Hepaticarum, 
but some of its most significant characters have since been empha- 
sized by Schiftner.* Considerable confusion was introduced by 
Taylor into the synonymy of the plant by the publication of Le- 
jeitnea domingensis. This species was based on specimens which 
he at first called L. linguaefolia, and material thus named was 
deposited in the Hooker herbarium. Subsequently he renamed 
the species L. domingensis and transferred the name L. linguaefolia 
to an entirely distinct plant collected by Richard on the island of 
St. Thomas. Both species were soon afterwards published in the 
same paper. Many years later, Spruce, basing his opinion on the 
specimens in the Hooker herbarium rather than on Taylor's de- 
scriptions, referred both L. domingensis and L. linguaefolia to his 
subgenus Acro-Lejennea, apparently considering them distinct from 
each other and also from Ptychocoleus poly carp us. Through the 
study of authentic material in the Lindenberg herbarium, Stephani t 
was able to reduce both species to synonymy, L. domingensis 
proving to be identical with Ptychocoleus polycarpus and the true 
L. linguaefolia from St. Thomas proving to be the same as Bra- 
chiolcjcunca cortkalis. Specimens in the Mitten herbarium, now in 
the possession of the New York Botanical Garden, amply confirm 
the conclusions of Stephani. 

The keels of the perianth in P. polycarpus scarcely extend 
below the middle and are exceedingly variable even on a single 
plant According to Schiffner, five or six keels are present, while 
Spru ce gives the n umber as four or five. These discrepancies are 

*Hedwigia 33: 181. 1894. See also Gottsche, Abhandl. nuturw. Verein. 
Bremen 7 r 350, 35^ 1882. 

fHedwigiaaS: 166, 167. 1889 ; 39: 22. 1890. 

Evans ; Hepaticae of Pukrto Rico 165 

doubtless due to differences of interpretation. When the antical 
surface is deeply grooved it produces the effect of two rounded 
keels, making the total number five or six, according to whether 
the confluent postical keels are counted as one or two* When the 
antical face is plane the number of keels would naturally be esti- 
mated as four. 

The two other species of Ptychocolats which have been reported 
from the West Indies are P. torulosus (Lehm. & Lindenb.) Trevis. 
and Lejatnea {Acrolejeuncd) atroviridis Spruce,* both of which are 
listed by Spruce from the island of St. Vincent, where they were 
collected by Elliott. The second of these species is known from 
no other locality, but P. torulosus is known also from Guiana and 
Brazil. In L. atroviridis the trigones of the leaves are absent or 
minute and the keels of the perianth are more or less roughened, 
so that it is possible that it ought to be referred to some other 
genus. P. torulosus, however, is a typical representative of Ptyeho- 
cohus. It may be at once distinguished from P. polycarpus by the 
blunt lobes and lobules of its bracts, by its truncate bracteole and 
by its seven- to nine-keeled perianth. It is distinct also m the 
lobules of its leaves, which bear from three to eight teeth along the 
free margin instead of one or two. 


According to Schiffner, the genus ArchiLjatnea contained thirty 
species in 1895, and perhaps a dozen new species have been pro- 
posed since he made his estimate. If, however, A. fordloidcs 
(Spruce) SchifTn., the first species described by Spruce, be selected 
as the type of the genus, certain species have been referred to it 
which can hardly be considered congeneric with this type species. 
This is the case, for example, with A. pseudocucullata Steph. {Lejat- 
nea holostipa Spruce), which the writer has already made the type of 
the genus Cyrtolejeunca^ It is true of A. xanthocarpa (Lehm. & 
Lindenb.) Steph. and its immediate allies, for the reception of which 
the genus Uucolejeunea Evans has recently been proposed. It is 
also true of A. conferta (Meissn.) Schiffn., which belongs to the 
genus Anoplolcjeunea and is even synonymous with i ts type species. 

- ■■ - ^m ^ 

^'" '■ ■ ■ 1 ■ - — ta*»_^ 

* Jour. Linn. Soc. BoL 30 : 335. /A 20. / j-10- 1894. 
t Bull. Torrey Club 30 : 55 2 « l 9°3* 

166 Evans : Hepaticae of Puerto Rico 

If these aberrant forms are removed, the species which are left 
will fully agree with the characters assigned to the genus 'by Spruce 
and by Schiffner. In the majority of cases the plants are robust 
and show a reddish or brownish pigmentation. A prostrate and 
irregularly branched caudex is first developed, which clings closely 
to the substratum (plate 8, figure 2). From this, secondary 
stems soon arise which are more or less free from the substratum 
and rarely form rhizoids ; they branch irregularly, many of the 
branches being subfloral innovations (figure i). Although the 
branches frequently bear smaller leaves than the stem, they are 
never truly microphyllous. 

The leaves are imbricated but not densely so, and the lobes 
spread obliquely or widely from the axis, being more or less fal- 
cate. They vary in outline from suborbicular to ligulate, the 
apices are rounded and the margins are entire. The lobules, in 
their most typical form, are rhomboidal in outline, the keel being 
straight or nearly so. In some cases about half of the lobule is 
appressed to the lobe, restricting the water-sac to a narrow linear 
space along the keel. In other cases the appressed portion is 
much narrower (figure 2) and may even be absent altogether, the 
free margin meeting the lobe at a right angle. Under these cir- 
cumstances the water-sac is relatively larger and the entire lobule 
may enter into its formation. The free margin is more or less 
curved and terminates in an apical tooth, which varies from suba- 
cute to long-acuminate ; the sinus beyond is long but usually 
shallow. The hyaline papilla is at the proximal base of the 
apical tooth and may be either marginal or slightly displaced to 
the inner surface of the lobule. Except for the apical tooth the 
margin is commonly entire. In certain species, however, a second 
tooth is sometimes developed, proximal in position to the papilla 
(figure 4). In some cases the secondary stems fail to develop 
normal lobules (figure i), and it becomes necessary to search for 
them on the prostrate caudex. The cells of the lobe have firm 
and pigmented walls and are usually a little convex. The trigones 
are large and conspicuous, and intermediate thickenings are fre- 
quent. The trigones are sometimes triangular but it is more 
usual for them to be triradiate with rounded rays (figure 3). 

The underleaves vary from orbicular to reniform. They are 

Evans: Hepaticae of Puerto Rico 167 

broad and rounded at the apex, entire along the margin, and vari- 
able at the base, being sometimes cuneate and sometimes rounded 
or subcordate. The line of insertion is slightly arched and there 
is sometimes a rudimentary radicelliferous disc at the base- Ex- 
cept on the caudex, however, rhizoids are very scantily de- 

In the majority of cases the inflorescence is dioicous, but a few 
autoicous species have been described. The female inflorescence 
is borne on a secondary stem or one of its leading branches and 
usually innovates on only one side. In a few species, however, 
two subfloral innovations are occasionally developed. An inno- 
vation spreads obliquely and frequently forms a second arche- 
gonium after bearing only one or two pairs of foliage leaves. 
When this procedure is repeated several times in succession, a 
cymose flower-cluster is the result, in which the flowers seem to 
be borne along the upper side of a floral axis. In rare cases an 
innovation is terminated by an antheridial spike. The bracts are 
unequally bifid and sharply complicate but are apparently never 
winged along the keel. The lobes spread obliquely and are rela- 
tively narrower than in the leaves, often showing a tendency to be 
sharp-pointed at the apex. The lobule is also narrow and varies at 
the apex from rounded to acute, The bracteole is free and ovate to 
obovate in outline, the apex showing all variations in different species 
from rounded or retuse to bidentate or bifid. In both bracts and 
bracteoles the margins are commonly entire. The perianth is 
oblong to obovate in outline, with a rounded or truncate apex and 
a short beak. It is more or less compressed with sharp lateral 
keels (figure 8). The antical surface commonly bears a short 
and low keel in the upper part, while the postical surface bears 
two sharp and confluent keels extending to below the middle. 
Both lateral and postical keels usually develop narrow and inter- 
rupted wings which are either irregularly sinuous or angular- 
dentate along the margin. In other respects the surface of the 
perianth is smooth. 

The male inflorescence is large and conspicuous, terminating a 
secondary stem or one of its leading branches ; in many cases, how- 
ler, it proliferates at the apex. The diandrous bracts are loosely 
imbricated and unequally bifid, both lobe and lobule being either 

168 Evans: Hepaticae of Puerto Rico 

rounded or very obtuse at the apex. The bracteoles extend the 
whole length of the spike and are much like the underleaves. 

Several genera of the Lejeuneae Holostipae are more or less 
closely allied to Archilejeunea. Certain species of Brachiolejeunea 
and Ptychocoleus, for example, develop a five-keeled perianth which 
is built up on a very similar plan. In Brachiolejeunea, however, 
the lobule is differently constructed and shows a larger number of 
marginal teeth, while in Ptychocoleus no subfloral innovations are 
present. Both of these genera, moreover, are composed of pros- 
trate species which show no distinction between caudex and 
secondary stems, and which are further characterized by the smooth 
and wingless keels of their perianths. In Mastigolejeunea the dis- 
tinction between caudex and secondary stems is well marked but 
the perianth is sharply trigonous with smooth keels. Spruce 
divided his subgenus Archi-Lejeunea into two sections : Monotro- 
pella, in which the keels of the perianth are rough and the sub- 
floral innovations usually occur singly ; and Dibrachiella, in which 
the keels are smooth and the innovations are sometimes borne in 
pairs. Schiffher accepts both of these sections, making them sub- 
genera of his genus Archilejeunea. Whether the slight differences 
just noted will ever be deemed sufficient to separate the groups 
generically is doubtful, since they share so many characters in 
common. It must be admitted, however, that Dibrachiella shows 
an especially close relationship to Brachiolejeunea and Ptycho- 

In its restricted sense Archilejeunea is almost exclusively trop- 
ical in its distribution. A number of species have been described 
from Africa and from the islands of the Pacific, but the highest 
development of the genus is attained in South America, where 
about half of the known representatives have been collected. At 
the present time no species are definitely known from Asia and 
only two species, both belonging to the subgenus Dibrachiella, 
have been reported from the West Indies. One of these is A. 
Auberiana (Mont.)Steph., originally described from Cuba, and the 
other is A. Cruegeri (Lindenb.) Schiffn., originally described from 
Trinidad. The distribution of these two species beyond the islands 
where they were first discovered is still imperfectly known. 
Stephani, to be sure, has reported A. Auberiana from Paramaribo 

Evans : Hepaticae of Puerto Rico 169 

and Para, * and Spruce has also reported A. Cruegeri from the 
latter locality, f Stephani's report is based on two specimens in the 
Lindenberg herbarium, one doubtfully referred to Lejeunea unci- 
loba Lindenb. and the other representing a portion of the original 
material of L. cydostipa Tayl. The writer has examined both of 
these specimens and would refer the first to L. (Arc/ii-Lejeum-a) 
florentissima Spruce rather than to A. Anberiana ; % the second is 
unfortunately so fragmentary that positive determination is hardly 
possible. With regard to A. Cruegeri the type material itself is 
scanty and poorly developed, making it difficult to obtain an ade- 
quate idea of the species. In view of these facts confirmatory 
evidence as to the occurrence of A. Anberiana and A. Cruegeri in 
South America is much to be desired. Although Archilcjeunea 
has not yet been reported from Puerto Rico, a single species, also 
belonging to the subgenus Dibrachiella, has been found on the 
island, namely : 

Archilejeunea viridissima (Lindenb.) 
Lejeunea viridissima Lindenb. in G. L. & N. byn. Hep. 320. 

Lejeunea {Archi- Lejeunea) viridissima Steph. Hedwigia 29: 17. 


Yellowish- or brownish-green, becoming darker with age, 
neither glossy nor glaucous, growing in depressed mats : caudex 
and secondary stems about O.14 mm. in diameter, the latter simple 
or sparingly subdivided, often with poorly developed lobules: 
leaves imbricated, the lobe widely spreading, slightly falcate, broadly 
ovate, 0.75 mm. long, 0.6 mm. wide, convex, and sometimes revo- 
lute at the broad and rounded apex and along the postical margin, 
antical margin straight or a little rounded at the base, then strongly 
outwardly curved to the apex ; lobule (when well developed) ovate, 
O.35 mm. long, 0.25 mm. wide, -the water-sac broad at the base 
and abruptly narrowed in the outer part, free margin sometimes 
revolute at the base, sometimes appressed to the lobe from base 
to apex, rounded, normally bearing two teeth in the outer part, 
the outer or apical a little longer than the other and often variously 
curved, the inner varying from acute to obtuse an d sometimes 

Hedwigia 29 : I5> 2I * 1 890. 
t Hep. Atnaz. et And* 97. 1884, 
% See Torreya 7 : 228. 1908. 

170 Evans : Hepaticae of Puerto Rico 

absent altogether, papilla marginal at base of apical tooth, sinus 
shallow, keel slightly arched, forming a very obtuse angle with the 
postical margin of the lobe ; cells of lobe plane or a little convex, 
averaging 12 fi at the margin, 24 x 18/i in the middle and 28 x 
18 /i at the base, trigones large, triradiate with rounded ends, 
separated from the oblong to circular intermediate thickenings by 
narrow pits : underleaves distant to loosely imbricated, appressed 
on the caudex but more or less squarrose on the secondary stems 
and sometimes revolute at the apex and along the sides, broadly 
orbicular-obovate, averaging 0.35 mm. in length and 0.4 mm. in 
width, cuneate at the base, rounded to subretuse at the apex: in- 
florescence polyoicous : 9 inflorescence terminating a secondary 
stem or one of its leading branches, innovating on one side or 
rarely on both, the innovations short and sterile or soon again 
floriferous, rarely terminating in an antheridial spike ; bracts erect- 
spreading, deeply and unequally bifid, complicate, the lobe oblong- 
obovate, 0.95 mm. long, 0.5 mm. wide, more or less convex and 
often revolute along the postical margin and at the apex, lobule 
ovate to ligulate, 0.4 mm. long, o. 1 5 mm. wide, rounded to subacute 
at the apex, keel very short ; bracteole obovate, averaging o. 7 5 X 0.4 
mm., rounded to retuse at the apex; perianth about half exserted, 
oblong-obovate in outline, cuneate toward the base, 1 .4 mm. long, 
0.75 mm. wide, antical keel low and broad, lateral and postical keels 
high and sharp, usually bearing narrow wings from one to three cells 
wide and variously sinuate orsubcrenulate on the margin : c? inflo- 
rescence terminating a short branch, apparently not proliferating ; 
bracts loosely imbricated, mostly in from three to six pairs, une- 
qually bifid, the lobe oblong to rounded at the apex, lobule similar 
but shorter and narrower, keel strongly arched ; antheridia in 
pairs ; bracteoles similar to the underleaves but smaller : capsule 
about 0.4 mm. in diameter, yellowish-brown ; spores greenish, 
irregular in form, about 16/* in short diameter, minutely verrucu- 
lose ; elaters 9,0 in diameter. (Plate 8, figures 1-8.) 

On a log. El Yunque, Evans (140). Type locality: Caracas, 
Venezuela (herb. Hampe). The species has also been found on 
the island of Great Bahama, Britton & Milhpmigh. The type 
material preserved in the Lindenberg herbarium agrees closely 
with the Puerto Rico and Bahamian specimens. 

Archdejeitnea viridissima is closely related to A. Auberiana. 
Through the kindness of M. Paul Hariot the writer has been sup- 
plied with a portion of the original material of the latter species 
from the Montagne herbarium. It shows an autoicous inflores- 
cence and agrees with A. viridissima in size, habit, and color. 

Evans: Hepaticae of Puerto Rico 171 

The lobules in both species are normally bidentate, and the 
perianths are five-carinate with smooth or nearly smooth keels. 
The lobule in A. Auberiana, however, is subrectangular in outline 
rather than ovate, the keel being nearly straight, and the marginal 
teeth tend to be larger and more pronounced than in A. viridis- 
sima. The leaf-cells also offer certain points of distinction, 
although they average about the same in size. In A. Aubcriana 
the local thickenings of the walls are rarely confluent, the trigones 
are mostly in the form of equilateral triangles with straight sides, 
and the intermediate thickenings are oval or circular in outline. 
In A. viridissima, on the other hand, the thickenings are more 
frequently confluent, the trigones show a strong tendency to be 
triradiate, with rounded and often constricted rays, and the inter- 
mediate thickenings show a similar tendency to be rectangular, 
with rounded ends. A. Aiiberiana is further characterized by the 
lobules of its perichaetial bracts ; these are much smaller than in 
A. viridissima and are usually adnate to the lobes throughout 
their entire length. Although the differences which have just 
been enumerated are slight, they seem to be constant. 

Another allied species is Lejennea florcntissima Spruce, which 
the writer would keep distinct from A. Auberiana. This species 
is more robust than A. viridissima, and the lobes of the leaves 
are more falcate ; the keel of the lobule is parallel with the free 
margin, but both are more or less curved; the trigones in the leaf- 
cells are similar to those in A, Aubcriana but are better developed, 
the sides of the triangles being convex rather than straight. The 
subfloral innovations in L. florcntissima are usually soon floriferous 
and the perianths are thus crowded together in a cymose cluster; 
in A. viridissima the arrangement of the perianths is much more 



The characters of the genus Leucolejeunta, recently segregated 
by the writer from Archil ejeune a, have already been so fully dis- 
cussed * that it hardly seems necessary to enumerate them here. 
Five species are definitely known at the present time, the generic 
type being L. clypeata (Schwein.) Evans, a widely distributed 
species in the eastern United States. The only representative of 

*See Torreya 7 ; 225-229, 1908. 

172 Evans : Hepaticae of Puerto Rico 

the genus which has been found in Puerto Rico is Z, xanthocarpa, 
although L. unciloha (Lindenb.) Evans is surely to be expected 
on the island, 

Leucolejeunea xanthocarpa (Lehm. & Lindenb.) Evans 

Jungermannia xanthocarpa Lehm. & Lindenb. in Lehmann, Pug, 

Plant. 5:8. 1832, 
Lejeunea xanthocarpa Lehm. & Lindenb.; G. L. & N. Syn. Hep. 

330, 1845. 
Lejeunea {Archi-Lejettned) xanthocarpa Pears, Christiana Vidensk.- 

Selsk, Forh. 1887 9 : 4. pL i.f. 14.-24, 
Archil ejettnea xanthocarpa Schiffn. Conspect. Hepat Archip. Indici 

316. 1898. 
Leucolejeunea xanthocarpa Evans, Torreya 7: 229. 1908. 

Pale glaucous-green, varying to bright-green and becoming 
yellowish or brownish upon drying, growing in depressed mats : 
stems 0.15 mm. in diameter, the branches obliquely to widely 
spreading, with smaller leaves than the stem but not microphyl- 
lous, sterile branches mostly simple or sparingly subdivided : leaves 
imbricated and usually densely so, the lobe (when flattened out) 
orbicular-ovate, 1 mm. long, 0,85 mm- wide, convex and strongly 
revolute at the broad and rounded apex and along the whole pos- 
tical margin, antical margin straight or slightly rounded at base, 
then outwardly curved to the apex ; lobule narrowly ovoid, 0.4 
mm. long t 0.2 mm. wide, gradually contracted in the outer half, 
the water-sac opening directly into the revolute portion of the lobe, 
keel straight or slightly arched continuous with revolute portion or 
slightly indented at junction, apex (not visible without flattening 
or dissection) varying from bluntly pointed to acuminate, in the 
latter case tipped with a row of from two to four cells, sinus straight 
or slightly iunulate, hyaline papilla mostly three to six cells from 
the apex ; cells of lobe plane or slightly convex, averaging 14/*** 
the margin, 20 p in the middle, and 28 x 20 (i at the base : under- 
leaves imbricated, plane (or a little convex from below), reniform, 
0.5 mm. long, 0,7 mm, wide, rounded to cordate at the base, 
rounded to vaguely retuse at the apex : 9 inflorescence borne on 
a more or less abbreviated branch, sometimes with only one pair 
of leaves below the involucre, innovating on one side ; bracts 
erect-spreading, the lobe falcate, convex, revolute at the rounded 
apex and along the postical margin, oblong-obovate, 1 mm, long, 
0.6 mm. wide, lobule oblong, 0,4 mm. long, 0.2 mm. wide, rounded 
to subacute at the apex, keel sometimes narrowly winged; brac- 
teole broadly obovate, 0.95 mm. long, 075 mm. wide, slightly 

Evans : Hepaticae of Puerto Rico 1 73 

retuse at the apex ; perianth almost hidden by the involucre, I mm. 
long, 0.7 mm. wide, narrowed toward the base, rounded to slightly 
retuse at the apex, beak 0.1-0.15 mm. long, keel roughened by 
projecting cells but destitute of wings or distinct teeth : o 71 in- 
florescence as described under the genus. (Plate 7, figures 

On trees. Near Cayey, Evans (76). The species is also 
known from the following islands of the West Indies: New 
Providence, Bahamas, A. E. Wight, E. G. Britton ; Jamaica, Un- 
derwood, Evans; Guadeloupe, Hn snot, Buss ; Dominica, Elliott; 
Martinique, Duss. On the American continent its range extends 
from Mexico to Brazil (the type locality). There are also records 
from Java and from Fernando Po, Mount Kilimanjaro and Cape 
Colony in Africa. A fragment of the type specimen in the Lin- 
denberg herbarium, collected by Beyrich in 1823 at New Freiburg, 
Brazil, agrees closely with the specimens here described. 

Apparently the closest allies of L. xantkocarpa tare L. unciloba, 
which has already been mentioned, and L. conchifolia Evans.* 
Both of these species have convex lobes and long teeth at the 
apices of the lobules. In L. unciloba, however, the apex of the 
lobe is only slightly revolute, and the apical tooth of the lobule is 
easily visible without dissection. In L. conchifolia the plants are 
less robust than in L. xantkocarpa, the lobes of the leaves spread 
more obliquely, the lobules are relatively larger, and their apical 
teeth are more like those of L. unciloba, being readily demonstrated 
without dissection. Z. xantkocarpa also bears a strong superficial 
resemblance to Pycnolejeuuca Sckivamckei (Steph.) Schiffii., which 
is of course at once distinguished by its bifid underleaves. 

The genus Anoplolcjeunca, as defined by Schifiner, contains the 
single species A. herpestica (Spruce) Schiffii. Apparently nothing 
has been written about this species except the rather brief descrip- 
tions of Spruce and Schiffner. The writer finds, however, upon 
examining a portion of the type material of A. herpestica^ kindly 

^^descriptions and figures of these two species see Evans, Mem. Torrey Club 
8: 125-129. pi. 16. /. it-zo ; pi f?. / '"V- ^02. L. unciloba is there called 
ArehiUjennea Stllowiana Steph., a species which the writer now considers synonymous 
with the much older Ltjeunea unciloba Lindenb. See also Torreya 7 I 229. 190S. 

174 Evans : Hepatic ae of Puerto Rlco 

sent by Mr. M. B. Slater, that it is quite indistinguishable from the 
widely distributed Lejeunea conferta Meissn. Although placed by 
recent writers in Archilejeunea, L. conferta cannot be considered 
congeneric with the typical members of this genus, and there seem 
to be excellent reasons for maintaining Anoplolejeuma as distinct 
Spruce compares it with his subgenus Plaiy-Lejetinea f but it is 
evidently much more closely related to the genus Leitcolejeunea* 

The plants grow in depressed mats and are either pale- or dark- 
green in color with neither glossiness nor pigmentation. The pros- 
trate stem is copiously and irregularly branched, some of the 
branches being similar to the stem while others are ascending and 
distinctly microphyllous (plate 8, figure 9). The normal leaves 
are imbricated, the convex lobe spreading widely and showing a 
rounded apex and entire margin. The leaf-cells are plane or 
nearly so, with small trigones and occasional intermediate thicken- 
ings (figure 1 6), and ocelli are sometimes present at the base of 
the lobe (figure 17). 

The lobule when normally developed is more complicated than 
in most of the other genera of the Lejeuneae* It is ovate-oblong 
in outline and forms a strongly inflated water-sac with an arched 
and convex keel (figure 9). From a relatively short base it is 
abruptly dilated and becomes suddenly contracted in the outer 
part. The dilated portion, bounded by the rounded and entire 
free margin, forms a volute with about one and a half turns (fig- 
ure 14). This comes into contact with the lobe, thus enclosing 
the water-sac, but the margin itself, together with a considerable 
extent of the lobule, lies free within the sac and can be distin- 
guished by careful focusing from below. The lobule develops no 
apical tooth, the free margin being directly continuous with the 
sinus; just at the junction of the two, however, a cell may be 
observed which is a little larger than its neighbors and which 
doubtless represents the terminal cell of the lobule, homologous 
with the apical tooth of other Lejeuncae (figure 18). At the 
beginning of its course the sinus is also strongly revolute but 
straightens out abruptly and passes into the postical margin of the 
lobe at the end of the keel. The hyaline papilla is marginal and 
arises from the terminal cell, usually at its junction with the first 
cell of the sinus ; it may therefore be regarded as distal in posi- 

Evans : Hepaticae of Puerto Rico 175 

tion. Unfortunately, the lobule is often poorly developed and 
fails to show some of the peculiarities which have just been 


The underleaves are distant and smaller than is usual among 
the Holostipae. They are obovate and entire, the apex varying 
from truncate to slightly retuse. When rhizoids are developed 
they usually spring from a distinct basal radicelliferous disc. 

The inflorescence is dioicous, and the male and female plants 
usually occur in separate mats. The female branch varies greatly 
in length and nearly always give rise to one or two subfloral inno- 
vations. In rare cases, however, the branch remains perfectly 
simple. The perichaetial bracts are a little larger than the leaves 
and their lobes are relatively narrower ; the lobules are rounded 
to subacute and the sharp keels are narrowly winged. Thebracteole 
is free and mostly obovate-oblong in outline, the apex varying from 
rounded to slightly retuse (figures 19, 21 ). The perianth is obo- 
void and normally shows five sharp keels (figure 23) and a short 
but distinct beak. The male inflorescence occupies a short branch 
or is terminal on a more or less elongated branch. The bracts 
are imbricated and diandrous, and the bracteoles are limited to the 
base of the spike. 

Anoplolejeunea conferta (Meissn.) 

Jungermannia conferta Meissn.; Sprengel in Linnaeus, Syst. Veg. 

ed. 16. 4: 325. 1827. 
Lejeunea subaurita Nees & Mont. Ann. Sci. Nat. Bot. II. 5 : 6a 

Lejeunea Miquclii Lehm. & Lindenb. in Lehmann, Pug. Plant. 7: 

15- 1 838. 

Lejeunea proteoides Lehm. & Lindenb, /. c. 7 : 19. 1838. 
Lejeunea conferta Meissn. ; G. L. & N. Syn. Hep. 333. 1845- 
Lejeunea involutiloba Mont Ann. Sci. Nat. Bot. IV. 5 : 3 50- 1856. 
Lejeunea (Anopio- Lejeunea) herpestica Spruce, Hep. Amaz. et And. 

129. 1884. 
Lejeunea {Platylejeunea) conferta Steph. Hedwigia 27 : 284. 

Lejeunea (Arc hi- Lejeunea) conferta Steph. /. c. 29 : 21. 1890. 
Anoplolejeunea herpestica Schiffn. in Engler & Prantl, Nat. Pflan- 

zenfam. I 3 : 131. 1895. 

176 Evans : Hepaticae of Puerto Rico 

Archilejeiuiea confcrta Schiffh. Conspect. Hepat Archip, Indici 

315. 1898. 

Plants becoming yellowish or brownish upon drying: stems 
0.17 mm. in diameter, branches obliquely to widely spreading: 
leaves imbricated, the lobe widely spreading, often revolute at the 
apex, falcate, broadly ovate, 1 mm. long, 0.85 mm. wide when of 
maximum size, often considerably smaller, arching across or a little 
beyond the axis, antical margin straight or slightly incurved near 
the base, then strongly outwardly curved to the apex, postical 
margin straight or somewhat curved, not revolute, forming an 
angle of 90 degrees or more with the strongly arched keel ; lobule 
0.35 mm. long, 0.2 mm. wide ; cells of lobe averaging 17 /i at the 
margin, 25 ft in the middle and 40 X 25 //at the base, trigones 
triangular with concave sides, intermediate thickenings sometimes 
confluent with the trigones ; ocelli (when present) mostly one to 
three, situated near the base of the lobe, measuring about 45 X 
32 fi: underleaves broadly orbicular-obovate, 0.35 mm. long, 
O.35-0.4 mm. wide, plane, cuneate and short-decurrent at the base : 
perichaetial bracts erect-spreading, the lobe obovate from a nar- 
row base, 1,2 mm. long, 0.7 mm. wide, convex and revolute at the 
apex, rounded to obtuse at the apex, lobule ovate-oblong, 0.75 
mm. long, 0.25 mm. wide; bracteole 0.4-0.75 mm. long, 0.25- 
0.4 mm. wide; perianth about two-fifths exserted at maturity, 
1-1,3 mm - long, 0.7-0.75 mm. in diameter, apex variable, usually 
truncate or subretuse but sometimes rounded or even subacute, 
keels variable, sometimes low and almost obsolete, sometimes dis- 
tinctly dilated in the upper part, when well developed showing a 
marginal row of slightly projecting cells with strongly thickened 
walls, postical surface of perianth sometimes developing one or two 
low and supplementary keels or folds : cT inflorescence apparently 
never proliferating ; bracts mostly in from two to seven pairs, 
closely imbricated, inflated, shortly bifid, the lobe strongly convex, 
rounded at the apex, antical margin straight or a little incurved, 
keel strongly arched, lobule obtuse to acute at the apex ; bracte- 
oles similar to the underleaves but smaller: mature sporophyte 
not seen. (Plate 8, figures 9-23.) 

On trees. Puerto Rico, without definite locality, SinUnis 
{rod). The writer has seen no specimens of A. confcrta from 
Puerto Rico but has collected it abundantly on the island of 
Jamaica, where it occurs at elevations of from 1,000 to 2,000 m. 
The species was originally collected in Peru but is now also known 
from Brazil, Bolivia, Venezuela, Trinidad, Colombia, and Mexico. 
Whether it occurs outside of tropical America is somewhat doubt- 

Evans: Hepaticae of Puerto Rico 177 

ful. The Synopsis Hepaticarum cites it from the islands of Luzon 
and Java, and SchifTner also quotes the latter locality. Both of 
these records, however, are based on old collections. 

The microphyllous branches of A. conferta (figure 9) are very 
characteristic, although they are sometimes scantily developed and 
may even be absent altogether. The lobes of their leaves are orbic- 
ular-ovate in outline and average about O.35 mm. in length ; they 
are suberect or spread obliquely from the axis instead of spread- 
ing widely as on normal stems and branches. The lobules are 
nearly spherical and measure only o. 1 5 mm. in diameter, in other 
respects agreeing with those on typical leaves. The underleaves, 
except for their small size, present no distinctive peculiarities. 
Under certain circumstances the lobules are poorly developed even 
on the stems and leading branches (ficure i 2) and thus give rise to 
forms which are not always easy to recognize. Usually a careful 
examination will reveal a few lobules of normal structure scattered 
among the others, and these will aid in the determination of such 
anomalous specimens. The characters upon which the varieties 
Miquelii and Liebmaniana of the Synopsis are based are apparently 
drawn from imperfectly formed lobules, and the propriety of 
attempting to maintain them is doubtful. 

In the specimens from Jamaica the lobes of the leaves are 
usually distinctly ocellate at the base except on poorly developed 
individuals. In all the South American specimens, however, which 
the writer has been able to examine, the ocelli are apparently ab- 
sent. If this difference should prove to be constant it might 
necessitate the separation of the West Indian plant as a distinct 
species. Unfortunately the South American specimens were all of 
old collections, and the detection of ocelli in plants of this charac- 
ter is sometimes very uncertain. Under the circumstances it seems 
best to include the Jamaican specimens with the others, a course 
which is advocated also by Stephani. 

A. conferta bears a rather strong superficial resemblance to 
Leucolejeunea clypeata, the two species agreeing in color and in 
many of the characters derived from leaf-cells, underleaves, pen- 
chaetial bracts, and perianths. They differ strikingly however, m 
the structure of the lobule, and L. clypeaia is also distinct m its 
inflorescence, which is normally autoicous, and in its lack ot micro- 

178 Evans: Hepaticae of Puerto Rico 

phyllous branches. There is little danger of confusing A. conferta 
with any other species of Leucolejeunea, in spite of the close re- 
lationship of the two genera. 

The present paper concludes the discussion of the Lejenmae 
Holostipae which are known to occur in Puerto Rico. Of the 
genera recognized by SchifTner the following four are apparently 
unrepresented on the island : Pty chant 1ms % Thysananthus, Peltole- 
jeunea and Dicranolejeunea. The last of these genera is abundant 
on Jamaica at rather high altitudes and may therefore be expected 
to have a wider distribution in the West Indies. There is little 
probability, however, that any of the others will be found there, 
although each is represented in South America by one or more 
species. Ptychanthus and Thysananthus, in fact, are essentially 
paleotropic in their distribution and attain their highest develop- 
ment in the East Indies and the neighboring parts of Asia. 

Yale University. 

Evans; Hepaticae of Puerto Rico 179 

Explanation of plates ft-£ 

As in the previous papers of this series the figures were drawn by the writer and 
prepared for publication by Miss Hyatt, 

Plate 6 

Brachi&hjeunea insularis Evans. Part of stem with branch, postical view, 
X 15- 2. Part of stem with base of branch, postical view, X ! S« 3- ^ eaf » antical 
view, X l S- 4? S* Perianths with involucres, the innovations dissected away, 
postical view, X 25. 6. Cells from middle of lobe, X 26 5- 7~9- Fr ^e margins of 
lobules, X 45* IO > ll - Apices of lobules, X 20 °- I2 * Marginal teeth of lobule, 
X 200. 13. Stem-leaf at base of branch, X *5- H- Basal auricle of underleaf, 
X 200. 15-17* Bracts and bracteole from one involucre, X f 5« 18-20. Bracts and 
bracteole from another involucre, X *5* 21, 22. Transverse sections of perianths, 
X35- Fig- 5 was drawn from specimen collected by Heller {44&3 a ) 5 Fi S s * *• 9» 
II, 18-20, from specimens collected by Howe (46J) ; the remaining figures from the 

type specimen* 

Plate 7 

Ptychccoleus polycarpm (Nees) Trevis. r. Part of stem with base of a branch, 
postical view, X f 5" 2 ' A P ex of f ema]e branch with perianth, postical view, X J 5- 
3, Part of stem, antical view, X<5* 4- Cells from middle of lobe, X 2 65« 5- Margin 
of lobule, the apex on right, X 2°°- 6 ^ S * Bracts and b ^cteole from the same invo- 
lucre, X J 5- 9- Subfloral leaf below involucre, X l S< l °- Bracteole from another 
specimen, X l 5* **« Transverse section of perianth, X 2 5- The Sg ures weTe a11 
drawn from specimens collected by Howe (1411* 1414}* 

Leucolejeunea xanthwarpa (Lehm. & Undent) Evans. 12. Part of sterile 
stem, postical view, X *5- *3- part o f P ]ant witl1 perianth, postical view, V 25* 14* 
Part of stem, antical view, X *5- *5- Leaf with lobule flattened out, X 15 l6 * Cel1s 
from middle of lobe, X265* 17. Cells from antical margin of lobe, X 200, 1 8, 19- 
Apices of lobules, X 200," 20-22. Bracts and bracteole from the same involucre, 
X25, 23. Transverse section of perianth, X 35* The figures were all drawn from 
specimens collected by the writer (76). 

Plate 8 

Architejtunea viridhiima (Lindenb.) Evans. I. Apex of female stem with peri- 
anth, postical view, X 2 5- 2 > part of caudex, postical view, X 2 5- 3- Cells from 
middle of lobe, X 26 5- 4- Margin of lobule, the apex on right, X 2< *>- 5i 6 - Bract 
and bracteole from the same involucre, X 2 5- 7* Bract from another involucre, X 2 5- 
8. Transverse section of perianth, X 2 5- The figures were all drawn from specimens 
collected by the writer (140)* 

AnoptoUjtunta conftrta ( Meissn. ) Evans. 9- P» rt of sterile stem with ^'W 1 " 
lous branches, postical view, X 25. 10. Female branch with perianth, postical view, 

X25. II. Apex of male branch, postical view, X »* I2 - Branch Wlth P °° t } Y de " 
veloped lobules, postical view, X 25. *3- Lobule, postical view, X 45- »4- Longi- 
tudinal section through lobe and lobule, X 45- *5- Mouth of lobule, diagrammatic. lb. 
Cells from middle of lobe, X 265. U- Cells from base of lobe, enclosing an ocellus. 
X200. 18. Margin of lobule, apical end on left, X200. IQ. Perianth with invo- 
lucre, postical view, X 35- 20, at. Bract and bracteole from a single involucre, 
X 25- 22. Perianth, postical view, X 25. ** Transverse section of penantb, X 35- 
The figures were all drawn from Jamaican specimens collected by the writer (jj, 272,